Quedius pallipes Lucas, 1849

Hansen, Aslak Kappel, Brunke, Adam, Simonsen, Thomas & Solodovnikov, Alexey, 2022, Revision of Quedius sensu stricto (Coleoptera: Staphylinidae), Acta Entomologica Musei Nationalis Pragae (Acta. Ent. Mus. Natl. Pragae) 62 (1), pp. 225-299 : 269-271

Quedius pallipes Lucas, 1849

( Figs 1 , 3 , 9C , 14D , 19 )

Quedius pallipes Lucas, 1849: 113 [Type locality: Algeria: environs de Philippeville]

Quedius leonhardi Bernhauer, 1914:67 [Type locality:Gargano], syn. rev.

Quedius levasseuri Coiffait, 1964: 293 [Type locality: Corse, Porto-Vecchio], syn. nov.

Quedius sardous Gridelli, 1924: 81 [Type locality: Sardegna: Cagliari, Assemini, Decimo, Golfo Aranci, Gonnesa], syn. rev.

Quedius simplicifrons Fairmaire, 1862: 580 [Type locality: Corse], syn. nov.

References. Fൺඎඏൾඅ (1869a):487, (1869b):151, (1902):115 (distribution; synonym of Q. molochinus ); Pඈඋඍൺ (1907): 130 (distribution; synonym of Q. molochinus ); MඣඊඎංGඇඈඇ (1937): 3 (distribution; subspecies of Q. molochinus ); Pൺඎඅංൺඇ (1941): 268 (larval characters; variety of Q. molochinus ); Lൺඌඍ (1952) (characters and distribution); KඈඋGൾ (1960): 53 (characters); (1962b):335, (1973): 275 (distribution); Cඈංൿൿൺංඍ (1961): 55 (characters); (1978):196 (distribution); Pൺඅආ (1963):142 (characters); SർHൾൾඋඉൾඅඍඓ (1965): 407; (1965b): 511 (distribution); Bඈඋൽඈඇං (1974): 12; (1976): 100 (characters and distribution); 1976b: 102 (characters); Cංർൾඋඈඇං & Zൺඇൾඍඍං (1995): 32 (distribution).

Type material examined. Quedius leonhardi : Hඈඅඈඍඒඉൾ J ( FMNH), labelled: “Monte Gargano 6.25.5.1907 legit. M. Hill Coll. O. Leonhard / Leonhardi Brh. Typus unique / Chicago NHMus M. Bernhauer Collection / In euparal / D.Drugmand det., 1994 Quedius (S.str.) molochinus (Grav.) / FMNHINS 3982385 FIELD MUSEUM Pinned / PHOTOGRAPHED S. Ware 2020 Emu catalog”.

Additional material examined. ALGERIA: Alger [Algiers], [36.71, 3.07], 2.-10.V.1972, leg. M. Dvořák (1 J MCZ); Bona [Annaba], [36.89, 7.76], leg. Meinert (1 ♀ NHMD); Mandoura, Boumerdes, [36.83, 3.66], 17.VI.1988, leg. Sama (1 J cSch). CHAD: Tibesti, Distr. Emi, Miski, [20.18, 17.94], 29.X.1885 (1 J NMW). FRANCE: Cඈඋඌൾ: Ajaccio, [41.93, 8.73] (1 J NHMD);Aleria, [42.11, 9.49], 1905, leg.O. Leonhard (1 J 1 ♀ SDEI). ITALY: Aspromonte, Samo, [38.07, 16.05], 250m, 12.III.2000, leg. F. Angelini (1 ♀ NHMD); Barletta, [41.32, 16.27], 14.V.1962, leg. J.T. Skovgaard (4 JJ 3 ♀♀ NHMD); Calaria, Locri, [38.25, 16.28], 3.V.1975, leg. R. Mourgua (1 ♀ cSch); Canale, [44.79, 7.99], leg. Paganetti (2 ♀♀ SDEI); Gargano, [41.71, 15.72], leg. L.S. Giovanni (1 ♀ NHMD); Lago S. Floriano, Toscana, [42.42, 11.35], 30.V.2002, leg. Bordoni (1 J 1 ♀ cBor); Lido di Roma, [41.73, 12.28], 17.-18.VI.1939 Palm (1 ♀ MZLU); Monti Aurunci, Itri, [41.31, 13.49], 25.IX.1973, Leg. M. Zampetti (1 ♀ cBor); Portonaccio, Lazio, [42.02, 12.39], 9.IX.1932, leg. G. Sacca (6 ♀♀ cBor); Puglia, Riserva Naturale Le Cesine, [40.35, 18.33], 11.VI.1999 leg. F. Angelini (2 ♀♀ NHMD); Roseto Capo Spulico, [39.98, 16.61], 10.XII.2000, leg. F. Angelini (1 ♀ NHMD). Sൺඋൽංඇංൺ: Assuni, [39.87, 8.94], leg. Krausse (1 ♀ SDEI); Cagliari, [39.21, 9.11], 29.I.1967, leg. S. Riese (3 ♀♀ cBor); Lago Omodeo, Oristano, [40.13, 8.92], 2.V.1979, leg.S. Riese (1J cBor); Oristano, [39.88, 8.58], 21.IV-7.V.1978, leg.S. Riese (1J 1♀ cBor); Ottana, [40.23, 9.04], 14.V.1976 (2 ♀♀ cBor); Sant’Antioco, [39.06, 8.45], IV.1984, leg. Bordoni (1 ♀ cBor); Sorgono, [40.02, 9.11], 1912 leg. Krausse (1 J SDEI); Stagni Cobras, Oristano, [39.91, 8.49], 19.IV.1971, leg. Visua (1 ♀ cBor). Sංർංඅඒ: Catania, [37.51, 15.09], IV.1965, leg. Fello (1 JcBor); Enna, [37.56, 14.28], 1.V.1990 leg. Roland (1 J NHMD); Gibilmanna, [37.98, 14.01], 800m 23.IV.1981, leg. T. Palm (1 ♀ MZLU); Marausa, [37.95, 12.5], 8.III.1982, leg.Alique (1 JcBor); Stracusa, [37.06, 15.28], 18.VII.1969, leg. Alraub (1 J 4 ♀♀ cBor). MOROCCO: Al Hakkama, ca. 18 km SE Tanger, 35.6377, -5.6841, fields, fallow, pastures, 100m, 26.II.2004, leg. D.W. Wrase (1 ♀ cSch); Bouznika, [33.81, -7.14], 12.III.1974 (1J cBor) Douyiet, [34.04, -5.12], 8.IV.1974 (1J 1♀ cBor); Fouarat, Rabat, [34.25, -6.52], 1932, leg.A.Théry (1 J NMW); 10 km N Mdiq, Tetouan, 35.7182, -5.3427, brackish wetland under plants in soil, 27.II.2004, leg. D.W. Wrase (1 J cSch); Mohammedia, Oued Mellah, Chefchaouen, 33.71, -7.33, stony slopes and bushes, 4.II.2003, leg. D.W. Wrase (1♀ cSch); Rif Mts. El-Malha, 19 km SE Bab-Taza, Chefchaouen, 34.9801, -5.0791, stony fallen land, ca. 440 m, 29.II.2004, leg. D.W.Wrase (1 ♀ cSch); 2 km W Tetouan, [35.55, -5.45], 16.II.1999, leg. Wrase (1 ♀ cSch). TUNISIA: El Besbassia, 10 km SW Hammamet,Sousee, [36.35, 10.52], 28.XI.-6.XII.1995, leg. D.W. Wrase (1 J cSch); El Kantaoui, 12 km NW Sousee, [35.89, 10.58], 25.-30.XI.1992, leg. D.W. Wrase (1 J cSch); Gabés, [33.88, 10.12], 30.V.- 5.6.1969, leg Palm (1 J MZLU).

Redescription. Measurements JJ (n = 4): HW = 1.64– 1.73 (1.69); HL = 1.47–1.51 (1.48); HL/HW 0.86–0.89 (0.88); PW = 2.16–2.31 (2.22); PL = 2.04–2.22 (2.11); PL/PW 0.93–0.96 (0.95); EW = 2.33–2.51 (2.39); EL = 2.16–2.36 (2.23); EL/EW 0.97–1.10 (1.06); EL/PL 0.97– 1.10 (1.06); PW/HW 1.47–1.55 (1.50); forebody length 5.69–6.00 (5.83). ♀♀ (n = 6): HW = 1.62–1.69 (1.67); HL = 1.36–1.47 (1.41); HL/HW 0.83–0.87 (0.84); PW = 2.13–2.27 (2.18); PL = 1.98–2.04 (2.03); PL/PW 0.90–0.96 (0.93); EW = 2.33–2.47 (2.41); EL = 2.13–2.29 (2.22); EL/EW 0.90–0.94 (0.92); EL/PL 1.07–1.12 (1.10); PW/ HW 1.45–1.62 (1.54); forebody length 5.49–5.73 (5.66).

Medium to large sized species; body black to light brown ( Fig. 9C ).

Head black, distinctly transverse, with eyes of medium size (EyL/TL: 1.76–2.20 (1.97)), microsculpture of transverse waves, no interocular setiferous punctures between anterior frontal punctures; antennae pale.

Thorax: pronotum paler dark brown to brown, slightly wider than long, clearly wider than head, with microsculpture of transverse waves, three punctures in dorsal row, sublateral row of pronotum with a cluster of one to three punctures situated slightly anterior of the anterior most puncture of the dorsal row. Scutellum with fine setiferous punctures and with wavy to meshed microsculpture; elytra usually light brown to brown, occasionally with paler suture and posterior margin, rarely completely black, relatively long, uniformly pubescent, slightly wider than long, of equal length or slightly longer than pronotum; legs pale, except inner tibia surface which is darkened.

Abdomen black to dark brown and slightly iridescent, tergites uniformly punctured, with slight iridescence.

Male. Aedeagus ( Fig. 14D ): paramere elongate, narrow, lanceolate with narrowly slightly asymmetrical apical portion exceeding apex of median lobe, with numerous sensory peg setae forming two subapical, lateral longitudinal groups connected apically below apex; median lobe with sides of apex folded into distinct teeth situated at medial part of fold. Internal sac with a pair of distal, very long medial sclerites, and with a pair of globular proximal sclerites.

Differential diagnosis. Quedius pallipes is distinguished from other Quedius s. str., except Q. hispanicus by a cluster of 1–3 additional setiferous punctures in the posterior part of the sublateral row of pronotum, which are very rarely missing. It can only be distinguished from Q. hispanicus by male genitalia characters; paramere slightly wider and less attenuate in middle portion, median lobe with sides of apex each folded into distinct tooth, tooth situated at about midlength of fold.

Synonymic notes. See under Quedius hispanicus . Based on the type locality, we here propose to use the name Q. pallipes for the Italian and North African species with the aedeagus type 2 ( Figs 14 , 19 ). Our examination of hundreds of specimens of the Q. pallipes -group across its full distribution range showed that specimens from Corse – which matched the original description of Q. simplicifrons – were conspecific with Q. pallipes . Therefore, Q. simplicifrons Fairmaire, 1862 , syn. nov. described from Corse, is here synonymized with Quedius pallipes Lucas, 1849 .

Quedius sardous Gridelli, 1924 from Sardegna was originally described as a variety of Q. picipennis [now Q. molochinus ]. Following Cඈൿൿൺංඍ (1961), Q. sardous was considered to be an infrasubspecific form of Q. simplicifrons . Here we move Quedius sardous Gridelli, 1924 syn. rev. to synonymy with Quedius pallipes Lucas, 1849 .

The original description of Quedius leonhardi Bernhauer, 1914 , a species described from Gargano in Italy and hitherto considered a synonym of Q. molochinus (e.g., Hൾඋආൺඇ 2001), matches Q. pallipes , except for the missing puncture in the sublateral row of pronotum, usually present in Q. pallipes . However, we have seen specimens of Q. pallipes (earlier named as Q. pallipoides , see below) with this puncture missing, while neither the described coloration of Q. leonhardi nor its distribution in central Italy match what we know about Q. molochinus . Thanks to S. Ware and M. Turcatel of the FMNH we were able to study images of the type of Q. leonhardi (see supplementary Fig. S4 ), which confirmed that Q. leonhardi fits the concept of Q. pallipes . Therefore, Q. leonhardi Bernhauer, 1914 , syn. rev. is here moved from synonymy with Q. molochinus ( Gravenhorst, 1806) to synonymy with Q. pallipes Lucas, 1849 .

Based on the original description of Cඈංൿൿൺංඍ (1964), and additional data by Bඈඋൽඈඇං (1976) and Cඈංൿൿൺංඍ (1978), Q. levasseuri, Coiffait 1964 , known only from the type series from Porto Vecchio on the island of Corse, we suggest that the male holotype have a teratological aedeagus. The shape of its aedeagus, not seen in any other specimen from this relatively well collected region, is the only unique trait of Q. levasseuri , because we observed its other presumably unique characters, the pitchy black elytra with narrow reddish margins, in both Q. hispanicus and Q. pallipes . Based on our hypothesis about the teratological aedeagus, we propose to synonymize Q. levasseuri , syn. nov. with Q. pallipes , which is also known from Corse.

Bionomics. Quedius pallipes is found at elevations from sea level to ca. 1000 m, in wet debris and under rocks, mainly near water sources. For example, based on examined specimen labels, it has been recorded in a brackish wetland under plants in soil, and on rocky slopes with bushes, both in interior Morocco. Based on records near coastal sites the species is most likely halotolerant.

Distribution. Quedius pallipes is distributed across northern Africa, the Italian Peninsula and the islands of Sicily, Sardinia and Corse ( Fig. 19 ). In northern Africa it is found in Morocco, Algeria and Tunisia, mainly in the coastal regions of these countries, or near water sources inland. A single record from Tibesti in Northern Chad is doubtful, as it is based on an unclear label, which may refer to the small settlement of Miski in the Emi Koussi District. In the Italian Peninsula it is found everywhere except its northern mountainous parts.

Figures

Fig. 1. Phylogenetic tree of Quedius s. str. based on preliminary target capture molecular dataset of Bඋඎඇĸൾ et al. (2021) and Hൺඇඌൾඇ et al. (in prep). Species groups shown in color. Q. altanai Hansen & Brunke, sp. nov. of the Q. molochinoides-group was not included in the dataset. Habitus sketch is of Q. fuliginosus (Gravenhorst, 1802). Lettering denotes species distribution within their respective groups.

Fig. 3. Phylogeny of Quedius s. str. based on Bayesian Inference (BI) and Maximum Likelihood (ML) analyses of COI barcodes (part 1). BI posterior probabilities (PP), ML ultrafast bootstrap support (UFB) and ML SH-aLRT test are shown. Nodes coloured as: green, PP ≥ 0.90, and either UFB ≥ 95 or SH-aLRT ≥ 80; yellow, one of PP ≥ 0.90, UFB ≥ 95 or SH-aLRT ≥ 80; red, PP <0.90, UFB <95 and SH-aLRT <80.

Fig. 9. Habitus of Quedius s. str. A – Q. hispanicus Bernhauer, 1898; B – Q. hispanicus (pale sutures variant); C – Q. pallipes Lucas, 1846; D – Q. laticollis (Gravenhorst, 1802) (black elytra variant); E – Q. laticollis (red elytra variant); F – Q. strenuus Casey, 1915.

Fig. 14. Aedeagus of Quedius s. str. A – Q. unicolor Kiesenwetter, 1847; B – Q. levasseuri Coiffait, 1964 (syn. of Q. pallipes Lucas, 1846); C – Q. hispanicus Bernhauer, 1898; D – Q. pallipes Lucas, 1846; E – Q. laticollis (Gravenhorst, 1802); F – Q. strenuus Casey, 1915. I – Parameral view of whole aedeagus, II – apices of paramere in antiparameral view with peg setae, III – lateral view of whole aedeagus, IV – apices of median lobe at an angle between parameral and lateral view.

Fig. 19. Distribution of Quedius levicollis Brullé, 1832 (top), Q. hispanicus Bernhauer, 1898 and Q. pallipes Lucas, 1846 (middle), and Q.balticus Korge, 1960 (bottom).

Fig. 4. Phylogeny of Quedius s. str. species based on Bayesian Inference (BI) and Maximum Likelihood (ML) COI barcodes (part 2). BI posterior probabilities (PP), ML ultrafast bootstrap support (UFB) and ML SH-aLRT test are shown. Nodes coloured as: green, PP ≥ 0.90, and either UFB ≥ 95 or SH-aLRT ≥ 80; yellow, one of PP ≥ 0.90, UFB ≥ 95 or SH-aLRT ≥ 80; red, PP <0.90, UFB <95 and SH-aLRT <80.